Spongesalso known as sea sponges), the members of the phylum Porifera[3] (/pəˈrɪfərəˌ pɔː-/ pər-IF-ər-ə, por-; meaning 'pore bearer'),[4] are a basal animal clade as a sister of the diploblasts.[5] They are multicellular organisms that have bodies full of pores and channels allowing water to circulate through them, consisting of jelly-like mesohyl sandwiched between two thin layers of cells.
Sponges have unspecialized cells that can transform into other types and that often migrate between the main cell layers and the mesohyl in the process. Sponges do not have complex nervous,[6] digestive or circulatory systems like humans. Instead, most rely on maintaining a constant water flow through their bodies to obtain food and oxygen and to remove wastes. Believed to be some of the most basal animals alive today, sponges were possibly the first to branch off the evolutionary tree from the last common ancestor of all animals, which would make them the sister group of all other animals.[5]
Sponges are similar to other animals in that they are multicellular, heterotrophic, lack cell walls and produce sperm cells. Unlike other animals, they lack true tissues[9] and organs.[10] Some of them are radially symmetrical, but most are asymmetrical. The shapes of their bodies are adapted for maximal efficiency of water flow through the central cavity, where the water deposits nutrients and then leaves through a hole called the osculum. The single-celled choanoflagellates resemble the choanocyte cells of sponges which are used to drive their water flow systems and capture most of their food. This along with phylogenetic studies of ribosomal molecules have been used as morphological evidence to suggest sponges are the sister group to the rest of animals.[11] A great majority are marine (salt-water) species, ranging in habitat from tidal zones to depths exceeding 8,800 m (5.5 mi), though there are freshwater species. All adult sponges are sessile, meaning that they attach to an underwater surface and remain fixed in place (i.e., do not travel). While in their larval stage of life, they are motile.
The few species of demosponge that have entirely soft fibrous skeletons with no hard elements have been used by humans over thousands of years for several purposes, including as padding and as cleaning tools. By the 1950s, though, these had been overfished so heavily that the industry almost collapsed, and most sponge-like materials are now synthetic. Sponges and their microscopic endosymbionts are now being researched as possible sources of medicines for treating a wide range of diseases. Dolphins have been observed using sponges as tools while foraging.[15]
Like cnidarians (jellyfish, etc.) and ctenophores (comb jellies), and unlike all other known metazoans, sponges' bodies consist of a non-living jelly-like mass (mesohyl) sandwiched between two main layers of cells.[19][20] Cnidarians and ctenophores have simple nervous systems, and their cell layers are bound by internal connections and by being mounted on a basement membrane (thin fibrous mat, also known as "basal lamina").[20] Sponges do not have a nervous system similar to that of vertebrates but may have one that is quite different.[6] Their middle jelly-like layers have large and varied populations of cells, and some types of cells in their outer layers may move into the middle layer and change their functions.[17]
A sponge's body is hollow and is held in shape by the mesohyl, a jelly-like substance made mainly of collagen and reinforced by a dense network of fibers also made of collagen. 18 distinct cell types have been identified.[23] The inner surface is covered with choanocytes, cells with cylindrical or conical collars surrounding one flagellum per choanocyte. The wave-like motion of the whip-like flagella drives water through the sponge's body. All sponges have ostia, channels leading to the interior through the mesohyl, and in most sponges these are controlled by tube-like porocytes that form closable inlet valves. Pinacocytes, plate-like cells, form a single-layered external skin over all other parts of the mesohyl that are not covered by choanocytes, and the pinacocytes also digest food particles that are too large to enter the ostia,[17][19] while those at the base of the animal are responsible for anchoring it.[19]
Glass sponges present a distinctive variation on this basic plan. Their spicules, which are made of silica, form a scaffolding-like framework between whose rods the living tissue is suspended like a cobweb that contains most of the cell types.[17] This tissue is a syncytium that in some ways behaves like many cells that share a single external membrane, and in others like a single cell with multiple nuclei.
Most sponges work rather like chimneys: they take in water at the bottom and eject it from the osculum at the top. Since ambient currents are faster at the top, the suction effect that they produce by Bernoulli's principle does some of the work for free. Sponges can control the water flow by various combinations of wholly or partially closing the osculum and ostia (the intake pores) and varying the beat of the flagella, and may shut it down if there is a lot of sand or silt in the water.[17]
The simplest body structure in sponges is a tube or vase shape known as "asconoid", but this severely limits the size of the animal. The body structure is characterized by a stalk-like spongocoel surrounded by a single layer of choanocytes. If it is simply scaled up, the ratio of its volume to surface area increases, because surface increases as the square of length or width while volume increases proportionally to the cube. The amount of tissue that needs food and oxygen is determined by the volume, but the pumping capacity that supplies food and oxygen depends on the area covered by choanocytes. Asconoid sponges seldom exceed 1 mm (0.039 in) in diameter.[17]
Some sponges overcome this limitation by adopting the "syconoid" structure, in which the body wall is pleated. The inner pockets of the pleats are lined with choanocytes, which connect to the outer pockets of the pleats by ostia. This increase in the number of choanocytes and hence in pumping capacity enables syconoid sponges to grow up to a few centimeters in diameter.
The "leuconoid" pattern boosts pumping capacity further by filling the interior almost completely with mesohyl that contains a network of chambers lined with choanocytes and connected to each other and to the water intakes and outlet by tubes. Leuconid sponges grow to over 1 m (3.3 ft) in diameter, and the fact that growth in any direction increases the number of choanocyte chambers enables them to take a wider range of forms, for example, "encrusting" sponges whose shapes follow those of the surfaces to which they attach. All freshwater and most shallow-water marine sponges have leuconid bodies. The networks of water passages in glass sponges are similar to the leuconid structure.[17]
In all three types of structure the cross-section area of the choanocyte-lined regions is much greater than that of the intake and outlet channels. This makes the flow slower near the choanocytes and thus makes it easier for them to trap food particles.[17] For example, in Leuconia, a small leuconoid sponge about 10 centimetres (3.9 in) tall and 1 centimetre (0.39 in) in diameter, water enters each of more than 80,000 intake canals at 6 cm per minute. However, because Leuconia has more than 2 million flagellated chambers whose combined diameter is much greater than that of the canals, water flow through chambers slows to 3.6 cm per hour, making it easy for choanocytes to capture food. All the water is expelled through a single osculum at about 8.5 cm per second, fast enough to carry waste products some distance away.[26]
In zoology a skeleton is any fairly rigid structure of an animal, irrespective of whether it has joints and irrespective of whether it is biomineralized. The mesohyl functions as an endoskeleton in most sponges, and is the only skeleton in soft sponges that encrust hard surfaces such as rocks. More commonly the mesohyl is stiffened by mineral spicules, by spongin fibers or both. Spicules, which are present in most but not all species,[27] may be made of silica or calcium carbonate, and vary in shape from simple rods to three-dimensional "stars" with up to six rays. Spicules are produced by sclerocyte cells,[17] and may be separate, connected by joints, or fused.[16]
Some sponges also secrete exoskeletons that lie completely outside their organic components. For example, sclerosponges ("hard sponges") have massive calcium carbonate exoskeletons over which the organic matter forms a thin layer with choanocyte chambers in pits in the mineral. These exoskeletons are secreted by the pinacocytes that form the animals' skins.[17]
It used to be claimed that glass sponges could live on nutrients dissolved in sea water and were very averse to silt.[29] However, a study in 2007 found no evidence of this and concluded that they extract bacteria and other micro-organisms from water very efficiently (about 79%) and process suspended sediment grains to extract such prey.[30] Collar bodies digest food and distribute it wrapped in vesicles that are transported by dynein "motor" molecules along bundles of microtubules that run throughout the syncytium.[17]
Sponges' cells absorb oxygen by diffusion from water into cells as water flows through body, into which carbon dioxide and other soluble waste products such as ammonia also diffuse. Archeocytes remove mineral particles that threaten to block the ostia, transport them through the mesohyl and generally dump them into the outgoing water current, although some species incorporate them into their skeletons.[17]
Most known carnivorous sponges have completely lost the water flow system and choanocytes. However, the genus Chondrocladia uses a highly modified water flow system to inflate balloon-like structures that are used for capturing prey.[33][36]
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